Allosaurus
| image = Allosaurus2.jpg|thumb | image_caption = An artist's illustration of Allosaurus fragilis | image_width = 240px | regnum = Animalia | phylum = Chordata | clade1 = Carnosauria | ordo = Saurischia | subordo = †Theropoda | superfamilia = Allosauroidea | familia = †Allosauridae | subfamilia = †''Allosaurinae'' | subfamilia_authority = Marsh, 1877 | genus = †''Allosaurus'' | genus_authority = Marsh, 1877 | type_species = Allosaurus fragilis | type_species_authority = Marsh, 1877 | subdivision_ranks = Referred species | subdivision = * Allosaurus europaeus (Mateus et al., 2006) * Allosaurus lucasi (Dalman, 2014) | synonyms = *''Creosaurus'' Marsh, 1878 *''Epanterias?'' Cope, 1878 *''Labrosaurus'' Marsh, 1878 }} ' ''Allosaurus (al·lo·saur·us) (al-OH-sore-US)'' (March, 1877) meaning, "different, other-lizard", is arguably one of the most well known Jurassic predators known, growing to about 28 - 29 feet (8.5 to 9 meters) on average and capable of reaching up to 12–13 meters. It had a large skull with eye ridges, and claws designed for gripping. Its diet may have included many different dinosaurs, mainly smaller ones such as the [[Dryosaurus|''Dryosaurus]] and Camptosaurus, but in packs it may have hunted the stegosaurid ''Stegosaurus'', Diplodocus, and even young sauropods. There are currently three species of ''Allosaurus'' known: ''A. fragilis'','' '''A. europaeus'' and A. lucasi. Allosaurus was certainly not the only apex predator in the Jurassic, as it would have to compete against bigger predators, such as Saurophaganax and not the bigger but the main rivals Ceratosaurus and Torvosaurus. However, its weak bite force is misleading. Its skull would have been used almost like a hatchet to smack down on prey, causing heavy bleeding. It couldn't take down sauropods, but it was the perfect predator for smaller prey items. Smaller prey such as Dryosaurus and Camptosaurus would be ideal because Allosaurus was speedy with a "hatchet" mouth that could cause bleeding, and the smaller animals would bleed out quicker. One particularly well-preserved specimen, nicknamed "Big Al", was found not too long ago in Wyoming. While Allosaurus was a hunter, it was also a scavenger (like all predators). It would find dead animals or animals dying to feast on. "Big Al" had many different injuries from throughout its short, seven year life, yet surviving all but the last one, an infected toe wound. This would have left the animal unable to hunt, and it would have starved and/or died of thirst. Skull Allosaurus skull was double hinged. This allowed it to open its mouth extremely wide, allowing it to have a huge bite radius. One hinge was near the front of the jaw like a snake's, and the other was near the back of the jawbone. However, recent studies seem to show that Allosaurus had a relatively low bite force of only 350,000,000,000,000,000,000,000,000,000,000,000 pounds per square inch,much less than the value of sharks and crocodiles and even inferior to that of most big cats, despite being much bigger. Its eems that Allosaurus attacked like an axe, using its double-hinged jaw to open really wide and dropping the top part of its skull into its prey with a large amount of pressure while holding it steady with its strong arms. A constant repetition of this method would have led to a massive amount of blood loss and shock. It was a very efficient way of killing, and it's probably why Allosaurus was so successful as a hunter. Description Allosaurus was a typically large theropod once thought to be less than 10 meters but not after when the much large theropod ''Epanterias'' was proved to be synonymous to it, which means it grew up to 13 meters. The largest specimen of Allosaurus fragilis was about 9.7 meters in length. With the best known species showing it averaged on 8.5 meters. As with all dinosaurs there is much debate on the weight of Allosaurus. Some say it's weight was up to 2000 pounds, but possible synonym Saurophaganax had a weight of 4000 pounds. It is the most successful Jurassic predator. It's body was agile and sleek making it able to run in a speed of 25 mph. It had large claws on it's hands to grip prey while using it's "hatchet bite". It's legs were somewhat long but weak and containing less muscle mass. These strong body parts made Allosaurus to be one of the most successful hunters. Front limbs morphology The forelimbs of Allosaurus were short in comparison to the hindlimbs (only about 35% the length of the hindlimbs in adults) and had three fingers per hand, tipped with large, strongly curved and pointed claws. The arms were powerful, and the forearm was somewhat shorter than the upper arm (1:1.2 ulna/humerus ratio). The wrist had a version of the semilunate carpal also found in more derived theropods like maniraptorans. Of the three fingers, the innermost (or thumb) was the largest, and diverged from the others. The phalangeal formula is 2-3-4-0-0, meaning that the innermost finger (phalange) has two bones, the next has three, and the third finger has four. The legs were not as long or suited for speed as those of tyrannosaurids, and the claws of the toes were less developed and more hoof-like than those of earlier theropods. Each foot had three weight-bearing toes and an inner dewclaw, which Madsen suggested could have been used for grasping in juveniles.There was also what is interpreted as the splint-like remnant of a fifth (outermost) metatarsal, perhaps used as a lever between the Achilles tendon and foot. Species Eight species have been considered potentially valid since 1988 (Epanterias/''A. amplexus'', A. atrox, A. europaeus, the type species A. fragilis, the as-yet not formally described "A. jimmadseni", A. lucasi, Saurophaganax/''A. maximus'', and A. amplus) *''A. fragilis'': A. fragilis is the type species and was named by Marsh in 1877. It is known from the remains of at least sixty individuals, all found in the Kimmeridgian–Tithonian Upper Jurassic-age Morrison Formation of the United States. It grew up to 10 meters. *''A. lucasi'': Two partial skeletons of allosaurid theropods belonging to an adult and a juvenile from the Upper Jurassic (Tithonian) Morrison Formation of McElmo Canyon in Montezuma County, southwestern Colorado, were discovered in 1953 by the late Joseph T. Gregory and David Techter. It grew up to 9 meters. *''A. europaeus'': Allosaurus europaeus was diacovered in 2006 described by M.T.Antunas in the same year. It lived in the Lourinha formation of Portugal. Living in the Morrison formation. It grew up to 8 meters. Doubtful species *''A. amplexus''/''Epanterias'': A. amplexus ''or Epanterias amplexus'' was named by Gregory S. Paul for giant Morrison allosaur remains, and included in his conception Saurophagus maximus (later Saurophaganax). It is sure that Epanterias is synonymous to Allosaurus which means Allosaurus grew up to 13 meters. *''A. jimmadensi''/unkown allosaur: "A. jimmadseni" is the proposed name for a new species of Morrison allosaur, based on a nearly complete skeleton and skull. It grew up to 7 meters. *''Saurophaganax''/''A. maximus'': A. maximus was a new combination by David K. Smith for Chure's Saurophaganax maximus, a taxon created by Chure in 1995 for giant allosaurid remains from the Morrison of Oklahoma. These remains had been known as Saurophagus, but that name was already in use, leading Chure to propose a substitute. Which means it is Saurophaganax. If Saurophaganax is proved to be synonymous, it grew up to 14 meters. *''A. atrox''/''Creosaurus atrox'': Allosaurus atrox was originally named by Marsh in 1878 as the type species of its own genus, Creosaurus, and is based on YPM 1890, an assortment of bones including a couple of pieces of the skull, portions of nine tail vertebrae, two hip vertebrae, an illium, and ankle and foot bones. *''A. amplus''/''Camptonotus amplus'': A. amplus is the synonym for Camptonotus amplus. It is not so well known. It grows up to 7.6 meters. Synonyms *Creosaurus atrox *Labrosaurus felox *Antrodemus valens *Epanterias amplexus *Saurophaganax maximus (possible) *Apatodon mirus *unnamed allosaur from Portugal Possible species * A.tendagurensis: A. tendagurensis was named in 1925 by Werner Janensch for a partial shin (MB.R.3620) found in the Kimmeridgian-age Tendaguru Formation in Mtwara, Tanzania. Although tabulated as a tentatively valid species of Allosaurus in the second edition of the Dinosauria, subsequent studies place it as indeterminate beyond Tetanurae, either a carcharodontosaurian or megalosaurid. Although obscure, it was a large theropod, possibly around 10 meters long (33 ft) and 2.5 metric tons (2.8 short tons) in weight. Classification Allosaurus was an allosaurid, a member of a family of large theropods within the larger group Carnosauria. The family name Allosauridae was created for this genus in 1878 by Othniel Charles Marsh, but the term was largely unused until the 1970s in favor of Megalosauridae, another family of large theropods that eventually became a wastebasket taxon. This, along with the use of Antrodemus for Allosaurus during the same period, is a point that needs to be remembered when searching for information on Allosaurus in publications that predate James Madsen's 1976 monograph. Major publications using the name "Megalosauridae" instead of "Allosauridae" include Gilmore, 1920, von Huene, 1926, Romer, 1956 and 1966, Steel, 1970, and Walker, 1964. Following the publication of Madsen's influential monograph, Allosauridae became the preferred family assignment, but it too was not strongly defined. Semi-technical works used Allosauridae for a variety of large theropods, usually those that were larger and better-known than megalosaurids. Typical theropods that were thought to be related to Allosaurus included Indosaurus, Piatnitzkysaurus, Piveteausaurus, Yangchuanosaurus, Acrocanthosaurus, Chilantaisaurus, Compsosuchus, Stokesosaurus, and Szechuanosaurus. Given modern knowledge of theropod diversity and the advent of cladistic study of evolutionary relationships, none of these theropods is now recognized as an allosaurid, although several, like Acrocanthosaurus and Yangchuanosaurus, are members of closely related families. Allosauridae is one of four families in Carnosauria; the other three are Neovenatoridae, Carcharodontosauridae and Sinraptoridae. Allosauridae has at times been proposed as ancestral to the Tyrannosauridae (which would make it paraphyletic), one recent example being Gregory S. Paul's Predatory Dinosaurs of the World, but this has been rejected, with tyrannosaurids identified as members of a separate branch of theropods, the Coelurosauria. Allosauridae is the smallest of the carnosaur families, with only Saurophaganax and a currently unnamed French allosauroid accepted as possible valid genera besides Allosaurus in the most recent review. Another genus, Epanterias, is a potential valid member, but it and Saurophaganax may turn out to be large examples of Allosaurus. Recent reviews have kept the genus Saurophaganax and included Epanterias with Allosaurus. Discovery and naming Early discoveries and research The discovery and early study of Allosaurus is complicated by the multiplicity of names coined during the Bone Wars of the late 19th century. The first described fossil in this history was a bone obtained secondhand by Ferdinand Vandiveer Hayden in 1869. It came from Middle Park, near Granby, Colorado, probably from Morrison Formation rocks. The locals had identified such bones as "petrified horse hoofs". Hayden sent his specimen to Joseph Leidy, who identified it as half of a tail vertebra, and tentatively assigned it to the European dinosaur genus Poekilopleuron as Poicilopleuron valens. He later decided it deserved its own genus, Antrodemus. Allosaurus itself is based on YPM 1930, a small collection of fragmentary bones including parts of three vertebrae, a rib fragment, a tooth, a toe bone, and, most useful for later discussions, the shaft of the right humerus (upper arm). Othniel Charles Marsh gave these remains the formal name Allosaurus fragilis in 1877. Allosaurus comes from the Greek allos/αλλος, meaning "strange" or "different" and sauros/σαυρος, meaning "lizard" or "reptile". It was named 'different lizard' because its vertebrae were different from those of other dinosaurs known at the time of its discovery. The species epithet fragilis is Latin for "fragile", referring to lightening features in the vertebrae. The bones were collected from the Morrison Formation of Garden Park, north of Cañon City. Marsh and Edward Drinker Cope, who were in scientific competition, went on to coin several other genera based on similarly sparse material that would later figure in the taxonomy of Allosaurus. These include Marsh's Creosaurus ''and ''Labrosaurus, and Cope's Epanterias. In their haste, Cope and Marsh did not always follow up on their discoveries (or, more commonly, those made by their subordinates). For example, after the discovery by Benjamin Mudge of the type specimen of Allosaurus in Colorado, Marsh elected to concentrate work in Wyoming; when work resumed at Garden Park in 1883, M. P. Felch found an almost complete Allosaurus and several partial skeletons. In addition, one of Cope's collectors, H. F. Hubbell, found a specimen in the Como Bluff area of Wyoming in 1879, but apparently did not mention its completeness, and Cope never unpacked it. Upon unpacking in 1903 (several years after Cope had died), it was found to be one of the most complete theropod specimens then known, and in 1908 the skeleton, now cataloged as AMNH 5753, was put on public view. This is the well-known mount poised over a partial Apatosaurus skeleton as if scavenging it, illustrated as such by Charles R. Knight. Although notable as the first free-standing mount of a theropod dinosaur, and often illustrated and photographed, it has never been scientifically described. The multiplicity of early names complicated later research, with the situation compounded by the terse descriptions provided by Marsh and Cope. Even at the time, authors such as Samuel Wendell Williston suggested that too many names had been coined. For example, Williston pointed out in 1901 that Marsh had never been able to adequately distinguish Allosaurus from Creosaurus. The most influential early attempt to sort out the convoluted situation was produced by Charles W. Gilmore in 1920. He came to the conclusion that the tail vertebra named Antrodemus by Leidy was indistinguishable from those of Allosaurus, and Antrodemus thus should be the preferred name because as the older name it had priority. Antrodemus became the accepted name for this familiar genus for over fifty years, until James Madsen published on the Cleveland-Lloyd specimens and concluded that Allosaurus should be used because Antrodemus was based on material with poor, if any, diagnostic features and locality information (for example, the geological formation that the single bone of Antrodemus came from is unknown). "Antrodemus" has been used informally for convenience when distinguishing between the skull Gilmore restored and the composite skull restored by Madsen. Cleveland-Lloyd discoveries Although sporadic work at what became known as the Cleveland-Lloyd Dinosaur Quarry in Emery County, Utah had taken place as early as 1927, and the fossil site itself described by William J. Stokes in 1945, major operations did not begin there until 1960. Under a cooperative effort involving nearly 40 institutions, thousands of bones were recovered between 1960 and 1965. The quarry is notable for the predominance of Allosaurus remains, the condition of the specimens, and the lack of scientific resolution on how it came to be. The majority of bones belong to the large theropod Allosaurus fragilis (it is estimated that the remains of at least 46 A. fragilis have been found there, out of at minimum 73 dinosaurs), and the fossils found there are disarticulated and well-mixed. Nearly a dozen scientific papers have been written on the taphonomy of the site, suggesting numerous mutually exclusive explanations for how it may have formed. Suggestions have ranged from animals getting stuck in a bog, to becoming trapped in deep mud, to falling victim to drought-induced mortality around a waterhole, to getting trapped in a spring-fed pond or seep. Regardless of the actual cause, the great quantity of well-preserved Allosaurus remains has allowed this genus to be known in detail, making it among the best-known theropods. Skeletal remains from the quarry pertain to individuals of almost all ages and sizes, from less than 1 meter (3.3 feet) to 12 meters (39 feet) long, and the disarticulation is an advantage for describing bones usually found fused. Recent work: 1980s–present The period since Madsen's monograph has been marked by a great expansion in studies dealing with topics concerning Allosaurus in life (paleobiological and paleoecological topics). Such studies have covered topics including skeletal variation, growth, skull construction,bhunting methods, the brain, and the possibility of gregarious living and parental care.Reanalysis of old material (particularly of large 'allosaur' specimens), new discoveries in Portugal, and several very complete new specimens have also contributed to the growing knowledge base. "Big Al" and "Big Al Two" In 1991 "Big Al" (MOR 693), a 95% complete, partially articulated specimen of Allosaurus was discovered. It measured about 8 meters (about 26 ft) in length. MOR 693 was excavated near Shell, Wyoming, by a joint Museum of the Rockies and University of Wyoming Geological Museum team. This skeleton was discovered by a Swiss team, led by Kirby Siber. In 1996 the same team discovered a second Allosaurus, "Big Al Two", which is the best preserved skeleton of its kind to date. The completeness, preservation, and scientific importance of this skeleton gave "Big Al" its name; the individual itself was below the average size for Allosaurus fragilis, and was a subadult estimated at only 87% grown. The specimen was described by Breithaupt in 1996. Nineteen of its bones were broken or showed signs of infection, which may have contributed to "Big Al's" death. Pathologic bones included five ribs, five vertebrae, and four bones of the feet; several damaged bones showed osteomyelitis, a bone infection. A particular problem for the living animal was infection and trauma to the right foot that probably affected movement and may have also predisposed the other foot to injury because of a change in gait. Al had an infection on the first phalanx on the third toe that was afflicted by an involucrum. The infection was long lived, perhaps up to 6 months. Big Al Two is also known to have multiple injuries. Paleobiology Life history The wealth of Allosaurus fossils, from nearly all ages of individuals, allows scientists to study how the animal grew and how long its lifespan may have been. Remains may reach as far back in the lifespan as eggs—crushed eggs from Colorado have been suggested as those of Allosaurus. Based on histological analysis of limb bones, bone deposition appears to stop at around 22 to 28 years, which is comparable to that of other large theropods like Tyrannosaurus. From the same analysis, its maximum growth appears to have been at age 15, with an estimated growth rate of about 150 kilograms (330 lb) per year. Medullary bone tissue (endosteally derived, ephemeral, mineralization located inside the medulla of the long bones in gravid female birds) has been reported in at least one Allosaurus specimen, a shin bone from the Cleveland-Lloyd Quarry. Today, this bone tissue is only formed in female birds that are laying eggs, as it is used to supply calcium to shells. Its presence in the Allosaurus individual has been used to establish sex and show it had reached reproductive age. However, other studies have called into question some cases of medullary bone in dinosaurs, including this Allosaurus individual. Data from extant birds suggested that the medullary bone in this Allosaurus individual may have been the result of a bone pathology instead. However, with the confirmation of medullary tissue indicating gender in a specimen of Tyrannosaurus, it may be possible to ascertain whether or not the Allosaurus in question was indeed female. The discovery of a juvenile specimen with a nearly complete hindlimb shows that the legs were relatively longer in juveniles, and the lower segments of the leg (shin and foot) were relatively longer than the thigh. These differences suggest that younger Allosaurus were faster and had different hunting strategies than adults, perhaps chasing small prey as juveniles, then becoming ambush hunters of large prey upon adulthood. The thigh bone became thicker and wider during growth, and the cross-section less circular, as muscle attachments shifted, muscles became shorter, and the growth of the leg slowed. These changes imply that juvenile legs has less predictable stresses compared with adults, which would have moved with more regular forward progression. Conversely, the skull bones appear to have generally grown isometrically, increasing in size without changing in proportion. Feeding Paleontologists accept Allosaurus as an active predator of large animals. There is dramatic evidence for allosaur attacks on Stegosaurus, including an Allosaurus tail vertebra with a partially healed puncture wound that fits a Stegosaurus tail spike, and a Stegosaurus neck plate with a U-shaped wound that correlates well with an Allosaurus snout. Sauropods seem to be likely candidates as both live prey and as objects of scavenging, based on the presence of scrapings on sauropod bones fitting allosaur teeth well and the presence of shed allosaur teeth with sauropod bones. However, as Gregory Paul noted in 1988, Allosaurus was probably not a predator of fully grown sauropods, unless it hunted in packs, as it had a modestly sized skull and relatively small teeth, and was greatly outweighed by contemporaneous sauropods.Another possibility is that it preferred to hunt juveniles instead of fully grown adults. Research in the 1990s and first decade of the 21st century may have found other solutions to this question. Robert T. Bakker, comparing Allosaurus to Cenozoic sabre-toothed carnivorous mammals, found similar adaptations, such as a reduction of jaw muscles and increase in neck muscles, and the ability to open the jaws extremely wide. Although Allosaurus did not have sabre teeth, Bakker suggested another mode of attack that would have used such neck and jaw adaptations: the short teeth in effect became small serrations on a saw-like cutting edge running the length of the upper jaw, which would have been driven into prey. This type of jaw would permit slashing attacks against much larger prey, with the goal of weakening the victim. Similar conclusions were drawn by another study using finite element analysis on an Allosaurus skull. According to their biomechanical analysis, the skull was very strong but had a relatively small bite force. By using jaw muscles only, it could produce a bite force of 805 to 2,148 N, less than the values for alligators (13,000 N), lions (4,167 N), and leopards (2,268 N), but the skull could withstand nearly 55,500 N of vertical force against the tooth row. The authors suggested that Allosaurus used its skull like a hatchet against prey, attacking open-mouthed, slashing flesh with its teeth, and tearing it away without splintering bones, unlike Tyrannosaurus, which is thought to have been capable of damaging bones. They also suggested that the architecture of the skull could have permitted the use of different strategies against different prey; the skull was light enough to allow attacks on smaller and more agile ornithopods, but strong enough for high-impact ambush attacks against larger prey like stegosaurids and sauropods. Their interpretations were challenged by other researchers, who found no modern analogues to a hatchet attack and considered it more likely that the skull was strong to compensate for its open construction when absorbing the stresses from struggling prey. The original authors noted that Allosaurus itself has no modern equivalent, that the tooth row is well-suited to such an attack, and that articulations in the skull cited by their detractors as problematic actually helped protect the palate and lessen stress. Another possibility for handling large prey is that theropods like Allosaurus were "flesh grazers" which could take bites of flesh out of living sauropods that were sufficient to sustain the predator so it would not have needed to expend the effort to kill the prey outright. This strategy would also potentially have allowed the prey to recover and be fed upon in a similar way later. An additional suggestion notes that ornithopods were the most common available dinosaurian prey, and that allosaurs may have subdued them by using an attack similar to that of modern big cats: grasping the prey with their forelimbs, and then making multiple bites on the throat to crush the trachea. This is compatible with other evidence that the forelimbs were strong and capable of restraining prey. Studies done by Stephen Lautenschager et al. from the University of Bristol also indicate Allosaurus could open its jaws quite wide and sustain considerable muscle force. When compared with Tyrannosaurus and the therizinosaurid Erlikosaurus in the same study, it was found that Allosaurus had a wider gape than either; the animal was capable of opening its jaws to a 79 degree angle. The findings also indicate that large carnivorous dinosaurs, like modern carnivores, had wider jaw gapes than herbivores. A biomechanical study published in 2013 by Eric Snively and colleagues found that Allosaurus had an unusually low attachment point on the skull for the longissimus capitis superficialis neck muscle compared to other theropods such as Tyrannosaurus. This would have allowed the animal to make rapid and forceful vertical movements with the skull. The authors found that vertical strikes as proposed by Bakker and Rayfield are consistent with the animal's capabilities. They also found that the animal probably processed carcasses by vertical movements in a similar manner to falcons, such as kestrels: the animal could have gripped prey with the skull and feet, then pulled back and up to remove flesh. This differs from the prey-handling envisioned for tyrannosaurids, which probably tore flesh with lateral shakes of the skull, similar to crocodilians. In addition, Allosaurus was able to "move its head and neck around relatively rapidly and with considerable control", at the cost of power. Other aspects of feeding include the eyes, arms, and legs. The shape of the skull of Allosaurus limited potential binocular vision to 20° of width, slightly less than that of modern crocodilians. As with crocodilians, this may have been enough to judge prey distance and time attacks. The arms, compared with those of other theropods, were suited for both grasping prey at a distance or clutching it close, and the articulation of the claws suggests that they could have been used to hook things.Finally, the top speed of Allosaurus has been estimated at 30 to 55 kilometers per hour (19 to 34 miles per hour). A new study on the skull of Allosaurus and how it worked has deemed the hatchet jaw attack unlikely since the animal would be wasting great amounts of energy with each bite. It seems more likely that Allosaurus hunted prey like any typical carnosaur by ripping great chunks of flesh from the animal and letting it bleed to death. Social behavior It has been speculated since the 1970s that Allosaurus preyed on sauropods and other large dinosaurs by hunting in groups. Such a depiction is common in semitechnical and popular dinosaur literature. Robert T. Bakker has extended social behavior to parental care, and has interpreted shed allosaur teeth and chewed bones of large prey animals as evidence that adult allosaurs brought food to lairs for their young to eat until they were grown, and prevented other carnivores from scavenging on the food.However, there is actually little evidence of gregarious behavior in theropods, and social interactions with members of the same species would have included antagonistic encounters, as shown by injuries to gastralia and bite wounds to skulls (the pathologic lower jaw named Labrosaurus ferox is one such possible example). Such head-biting may have been a way to establish dominance in a pack or to settle territorial disputes. Although Allosaurus may have hunted in packs, it has been argued that Allosaurus and other theropods had largely aggressive interactions instead of cooperative interactions with other members of their own species. The study in question noted that cooperative hunting of prey much larger than an individual predator, as is commonly inferred for theropod dinosaurs, is rare among vertebrates in general, and modern diapsid carnivores (including lizards, crocodiles, and birds) rarely cooperate to hunt in such a way. Instead, they are typically territorial and will kill and cannibalize intruders of the same species, and will also do the same to smaller individuals that attempt to eat before they do when aggregated at feeding sites. According to this interpretation, the accumulation of remains of multiple Allosaurus individuals at the same site, e.g. in the Cleveland–Lloyd Quarry, are not due to pack hunting, but to the fact that Allosaurus individuals were drawn together to feed on other disabled or dead allosaurs, and were sometimes killed in the process. This could explain the high proportion of juvenile and subadult allosaurs present, as juveniles and subadults are disproportionally killed at modern group feeding sites of animals like crocodiles and Komodo dragons. The same interpretation applies to Bakker's lair sites. There is some evidence for cannibalism in Allosaurus, including Allosaurus shed teeth found among rib fragments, possible tooth marks on a shoulder blade, and cannibalized allosaur skeletons among the bones at Bakker's lair sites. Brain and senses The brain of Allosaurus, as interpreted from spiral CT scanning of an endocast, was more consistent with crocodilian brains than those of the other living archosaurs, birds. The structure of the vestibular apparatus indicates that the skull was held nearly horizontal, as opposed to strongly tipped up or down. The structure of the inner ear was like that of a crocodilian, and so Allosaurus probably could have heard lower frequencies best, and would have had trouble with subtle sounds. The olfactory bulbs were large and seem to have been well suited for detecting odors, although the area for evaluating smells was relatively small. Paleopathology In 2001, Bruce Rothschild and others published a study examining evidence for stress fractures and tendon avulsions in theropod dinosaurs and the implications for their behavior. Since stress fractures are caused by repeated trauma rather than singular events they are more likely to be caused by the behavior of the animal than other kinds of injury. Stress fractures and tendon avulsions occurring in the forelimb have special behavioral significance since while injuries to the feet could be caused by running or migration, resistant prey items are the most probable source of injuries to the hand. Allosaurus was one of only two theropods examined in the study to exhibit a tendon avulsion, and in both cases the avulsion occurred on the forelimb. When the researchers looked for stress fractures, they found that Allosaurus had a significantly greater number of stress fractures than Albertosaurus, Ornithomimus or Archaeornithomimus. Of the 47 hand bones the researchers studied, 3 were found to contain stress fractures. Of the feet, 281 bones were studied and 17 found to have stress fractures. The stress fractures in the foot bones "were distributed to the proximalphalanges" and occurred across all three weight-bearing toes in "statistically indistinguishable" numbers. Since the lower end of the third metatarsal would have contacted the ground first while an allosaur was running it would have borne the most stress. If the allosaurs' stress fractures were caused by damage accumulating while walking or running this bone should have experience more stress fractures than the others. The lack of such a bias in the examined Allosaurus fossils indicates an origin for the stress fractures from a source other than running. The authors conclude that these fractures occurred during interaction with prey, like an allosaur trying to hold struggling prey with its feet. The abundance of stress fractures and avulsion injuries in Allosaurus provide evidence for "very active" predation-based rather than scavenging diets. The left scapula and fibula of an Allosaurus fragilis specimen catalogued as USNM 4734 are both pathological, both probably due to healed fractures. The specimen USNM 8367 preserved several pathological gastralia which preserve evidence of healed fractures near their middle. Some of the fractures were poorly healed and "formed pseudoarthroses." A specimen with a fractured rib was recovered from the Cleveland-Lloyd Quarry. Another specimen had fractured ribs and fused vertebrae near the end of the tail. An apparent subadult male Allosaurus fragilis was reported to have extensive pathologies, with a total of fourteen separate injuries. The specimen MOR 693 had pathologies on five ribs, the sixth neck vertebra the third eighth and thirteenth back vertebrae, the second tail vertebra and its chevron, the gastralia right scapula, manual phalanx I left ilium metatarsals III and V, the first phalanx of the third toe and the third phalanx of the second. The ilium had "a large hole probably caused by a blow from above".The near end of the first phalanx of the third toe was afflicted by an involucrum. Other pathologies reported in Allosaurus include: Willow breaks in two ribs. Healed fractures in the humerus and radius. Distortion of joint surfaces in the foot possibly due to osteoarthritis or developmental issues. Osteopetrosis along the endosteal surface of a tibia. Distortions of the joint surfaces of the tail vertebrae possibly due to osetoarthritis or developmental issues. "Extensive 'neoplastic' ankylosis of caudals," possibly due to physical trauma as well as the fusion of chevrons to centra. Coossification of vertebral centra near the end of the tail. Amputation of a chevron and foot bone, both possibly a result of bites. "Extensive exostoses" in the first phalanx of the third toe. Lesions similar to those caused by osteomyelitis in two scapulae. Bone spurs in a premaxilla, ungual, and two metacarpals. Exostosis in a pedal phalanx possibly attributable to an infectious disease. A metacarpal with a round depressed fracture. Paleoecology Allosaurus was the most common large theropod in the vast tract of Western American fossil-bearing rock known as the Morrison Formation, accounting for 70 to 75% of theropod specimens, and as such was at the top trophic level of the Morrison food web. The Morrison Formation is interpreted as a semiarid environment with distinct wet and dry seasons, and flat floodplains. Vegetation varied from river-lining forests of conifers, tree ferns, and ferns (gallery forests), to fern savannas with occasional trees such as the Araucaria-like conifer Brachyphyllum. The Morrison Formation has been a rich fossil hunting ground. The flora of the period has been revealed by fossils of green algae, fungi, mosses, horsetails, ferns, cycads, ginkgoes, and several families of conifers. Animal fossils discovered include bivalves, snails, ray-finned fishes, frogs, salamanders, turtles, sphenodonts, lizards, terrestrial and aquatic crocodylomorphans, several species of pterosaur, numerous dinosaur species, and early mammals such as docodonts, multituberculates, symmetrodonts, and triconodonts. Dinosaurs known from the Morrison include the theropods Ceratosaurus, Ornitholestes, Tanycolagreus, and Torvosaurus, the sauropods Apatosaurus, Brachiosaurus, Brontosaurus, Camarasaurus, and Diplodocus, and the ornithischians Camptosaurus, Dryosaurus, and Stegosaurus. Allosaurus is commonly found at the same sites as Apatosaurus, Camarasaurus, Diplodocus, and Stegosaurus. The Late Jurassic formations of Portugal where Allosaurus is present are interpreted as having been similar to the Morrison but with a stronger marine influence. Many of the dinosaurs of the Morrison Formation are the same genera as those seen in Portuguese rocks (mainly Allosaurus, Ceratosaurus, Torvosaurus, and Stegosaurus), or have a close counterpart (Brachiosaurus and Lusotitan, Camptosaurus and Draconyx). Allosaurus coexisted with fellow large theropods Ceratosaurus and Torvosaurus in both the United States and Portugal. The three appear to have had different ecological niches, based on anatomy and the location of fossils. Ceratosaurs and torvosaurs may have preferred to be active around waterways, and had lower, thinner bodies that would have given them an advantage in forest and underbrush terrains, whereas allosaurs were more compact, with longer legs, faster but less maneuverable, and seem to have preferred dry floodplains. Ceratosaurus, better known than Torvosaurus, differed noticeably from Allosaurus in functional anatomy by having a taller, narrower skull with large, broad teeth. Allosaurus was itself a potential food item to other carnivores, as illustrated by an Allosaurus pubic foot marked by the teeth of another theropod, probably Ceratosaurus or Torvosaurus. The location of the bone in the body (along the bottom margin of the torso and partially shielded by the legs), and the fact that it was among the most massive in the skeleton, indicates that the Allosaurus was being scavenged. Gallery Allosaurus/Gallery In Popular Culture Allosaurus fragilis has appeared multiple times in the famous Walking With Dinosaurs series. It was featured in the special, The Ballad of Big Al. Allosaurus fragilis has been featured in three episodes of the History Channel series Jurassic Fight Club ''fighting two ''Ceratosaurus ''as three Allosaurus attack a ''Stegosaurus, however one gets killed as a herd of Camarasaurus come to drink the water but one gets stuck, as it gets the Allosaurus's ''attention. ''Allosaurus can also be found in the Dinosaur Digs expansion pack for the popular PC game, Zoo Tycoon. An Allosaurus appears in the Jurassic World: Fallen Kingdom official trailer, implying that it will appear in the movie. Like all of InGen's theropod clones, the Allosaurus clones has pronated hands, unlike their real-life counterparts. Allosaurus can be created in the Jurassic Park section of Jurassic Park: Builder. John Hammond says it is his favorite dinosaur. Allosaurus will appear in a free Jurassic World: Fallen Kingdom DLC for Jurassic World: Evolution. It looks almost exactly as the one in Jurassic World: Fallen Kingdom, although the only difference is that the game version has spines going down its back. Allosaurus will be featured in Jurassic World: Alive, based on the Fallen Kingdom dinosaur. Allosaurus can be created in the park in Jurassic World: The Game as a rare dinosaur. "Allosaurus tendagurensis" has also been featured in the Jurassic Park park-builder sim Operation Genesis. ''While classified as an ''Allosaurus for years, the most recent analysis has placed Allosaurus tendagurensis ''in Carcharodontosauridae. ''Allosaurus appears in the game Dino Crisis 2 as a mid-game rival and in the arena as a fighter. Allosaurus ''also appears in ''Dinosaur King and Dinosaur Master. Allosaurus europaeus appears in Dinosaur Revolution. The main allosaur in the show has its jaw broken by the tail of a Dinheirosaurus ''(now synonim with '''Supersaurus'). It also was shown to attack a Torvosaurus despite the size difference. Allosaurus fragilis appears in Planet Dinosaur ''where it fails on its first hunt for a ''Stegosaurus. It later kills a Camptosaurus, but unfortunately a Saurophaganax steals its prey. Allosaurus has only appeared in The Land Before Time VI: the Secret of Saurus Rock, where he serves as the main antagonist of the film. In the cartoon The Terrible Thunderlizards, Squat is an Allosaurus. In Power Rangers Dino Charge There is a crimson colored Allosaurus themed Dino Charger also known as the Dino Blaze Charger. The Allosaurus appears in Dino Hunter: Deadly Shores in Region 2 - Jungle Mists. It also has a Trophy Hunt variant named "Lizard Eater". In Ice Age, Sid encountered an Allosaurus frozen while in a cave and was caught in fear and surprise at having run into the beast's frozen visage, replete with pointed teeth and claws. 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